PSYCHE AND WESTERN SYRIA PART II. In 1906 BSrner described Cyphoderus assimilis from THE COLLEMBOLA OF LEBANON FAMILIES CYPHODERIDAE AND ONCOPODURIDAE

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PSYCHE Vol. 64 September, 1957 No. 3 THE COLLEMBOLA OF LEBANON AND WESTERN SYRIA PART II. FAMILIES CYPHODERIDAE AND ONCOPODURIDAE BY K. CHRISTIANSEN Grinnell College, Grinnell, Iowa In 1906 BSrner described Cyphoderus assimilis from material partly collected in Egypt. Since that time the family Cyphoderidae has been well represented in the collections from the region until at the present moment nine species have been recorded from Lebanon, Palestine, Egypt, and Iraq. Four.of the nine species mentioned above were again recovered in the course of this research and two new species were uncovered. The relatively poor recovery of previously located species along with the extremely spotty nature of the distribution of the collections appears to indicate a small sampling of the total population of this family present in the area. The family Oncopoduridae has not previously been reported from the. area. The single new species described herein does not give any new insight into the biogeography of the region. Genus Cyphoderus Delamare-Deboutteville (1948) in his excellent work upon Termitophilous and Myrmecophilous Collembola divided the old genus Cyphoderus into a number of genera. Since two of the species here examined indicated a clear need for the re-examination of these genera, I have con- Published with the aid of a grant from the Museum of Comparative Zoology at Harvard College. Part I (General Considerations and the Family Onychiuridae) was published in Psyche 6 (4): 119-133, 1956. 77

78 Psyche [September sidered his g.enera as subgenera (Cyphoda and Serroderus) throughout, merely to avoid con2usion. Three species had been previously recorded from Lebanon. In the. present work two o2 these were re-discovered, and species previously recorded rom Egypt and Palestine were found. Along with one new species, this brings the total known from the area to. six species. Cyphoderus genneserae Carpenter Cyphoderus genneserae Carpenter, 1913, Jour. Proc. Asiat. Soc. Bengal 9:215 This species is fairly widespread throughout the southwestern Syrian Region. Handschin and Carpenter reported it from Palestine and the variation described by Handschin as "aethiopica" has been found in several regions in Central Africa. I have recovered the speci.es from two localities" Litani River near the mouth on August 23, 1953 (Asfour and Salaymeh Coll.) and Antelias Stream Bank on November 9, 1952. Both.of these l.ca!ities are in the coastal plain o2 Lebanon and all.o2 the specimens (about 25) agree with the descriptions and figures shown in Handschin, except in one startling respect: The presence o2 only two fringed scales in the inner row of dental scales. This characteristic places the species in the genus Serroderus of Delamare, although it lacks any hint.o the gradation between scale and seta structure described as characteristic of the genus. The species of this genus which it most resembles is S. sublimis and this species also lacks the above mentioned characteristics. In spite of the difficulty of separating the specimens at hand from the last mentioned species I 2eel that the exact detail for detail identity of the present orm with the described gennesera.e leaves little doubt that the existence of only a single pair of internal scales is a local variation. This, o. course, points up the necessity.of revising the limitations of the genus Serroderus, but this is beyond the scope of the present paper. The only other modification.of Handschin s description is that in a few specimens the tenent hair can be seen to have a minutely clavate tip, visible only under the highest magnifi-

Christiansen Collembola of Lebanon 79 cation. In both populations the aethiopica type of mucronal form was present in a.ddition to. the normal type. Cyphoderus (Cyphoda) grassei Cassagnau & Delamare-Deboutteville Cyphoda grassei, Cassagnu & Delamare, 1951, Biospeologica 75: 384-385. This species was described by Cassagnau and Delamare 2rom a single specimen. The relative size the mucro and dens (1-10) was taken as a. characteristic o the species and upon this basis it was placed in the genus Cyphoda. I have several series of specimens which I have identified as this species. In one of these I have five specimens. The r.elative sizes.of the mucro and dens varying as follows" 1-10, 1-9, 1-8.5, 1-7.5, 1-7. In a second series.of three specimens the sizes are: 1-10, 1-6, and 1-5.5. In still a third series both specimens have a mucronal dent,es ratio of 1-5.5. This presents a problem since the sole basis for separation o members.of the genus Cyphoda is the relativ.e mucr.onal length. Whatever the eventual decision concerning this genus, the species grassei can no longer be considered part of the genus. In addition to. this variation in mucr.o the species shows another peculiarity in that the tunica is absent in a few specimens, particularly young forms. In over 95% of the specimens seen, a clear tunica is present on the ungues. Perhaps the most unique characteristic of the group is the shape.of the mucro. This is particularly noticeable upon specimens having a relatively large mucro, but can be seen upon all. The mucro has in addition to the apical hook, a very shallow anteapical tooth. This condition is always the same and places it directly.on a line between the two groups bidenticulati and inermes. Distribution" Hammana, Lebanon, August 28, 1952 and July 28, 1957. Vic. East Base Quornet es Sauda Lebanon, el,ev. 2200 meters, July 1, 1953; Vic. Turkish Border Latakia, Syria, August 2, 1953, lat. 35 50" long. 36 01"; Dahr el Ain, Lebanon, October 10, 1951. Cyphoderus (Serroderus) spinatus n. sp. Plate 6, figures 9-11, Plate 7, figure 8 Body oval, head bluntly oval with well developed mouth-

80 Psyche [September parts. White without eyes or any indication of pigment. Antennae our-segmented with first and second segments subcylindrical, third segment short and swollen apically, and fourth segment fusiform. Clothing of fourth segment of numerous setae some curved acuminate and ciliate, others smooth blunt and sharply curved. Both varieties decrease in size apically. The remainder of antennal segments are similarly clothed except that ciliate seta.e become relatively larger and straighter, and the smooth setae are largely limited to the ventral surfaces. The third ant.ennal sense organ of tw.o blunt ovoid stalked knobs and a pair of small slender, sharply angled blunt setae. The body well covered with fine scales. Large ciliate acuminate setae present.on the ventral surface of the animal and posterior two segments. Body with scattered short to 10ng slender bothriotrichia. Legs with numerous short acuminate setae. Tenent hair stout and pointed on all legs. Unguis stout, sharply acuminate with paired basal prominent teeth, two small but definite median teeth. Apical-most tooth at about mid-level of unguis. Manubrium with normal clothing, dens with acuminate ciliate setae, ciliate and smooth scales and spines (see Plate 6, fig. 10). Outer row of dorsal fringed scales six EXPLANATION OF I)LATE 6 Figures 1-50ncopodura ambigua n. sp. 1. Claw, paratype, approx.!000 x. 2. Habitus, setae and scales omitted, holotype, approx. 100 x. 3. Left postantennal organ, paratype, approx. 1000 x. 4. Third antennal sense organ, paratype, approx. 2000 x. 5. Dens and mucro, holotype. approx. 2000 x. Figures 6-8 Troglopedetes canis n. sp. 6. Detail of clothing of antenna, with different types of setae mentioned in description labelled. Small lower diagram shows whole fourth segment with expanded part hatched, detail holotype, approx. 1000 x. 7. Front claw, holotype, approx. 1000 x. 8. End of dens and mucro, paratype, approx. 350 x. Figures 9-11 Cyphoderus spinatus n. sp. 9. Clasp organ at base, of dens, paratype, approx. 1000 x. 10. Mucro and dens, holotype, approx. 400 x. 11. Hind foot, paratype, approx. 400 x. Figure 12. Cyphoderus assimilis BSrner. Mucro and dens of specimen from Syria. approx. 400 x.

]:)SYCI-IE, 1957 VOL. 64, PLATE 6 C]tRISTIANSEN CYPI-tODERUS

82 Psyche [eptember in number,,distalmost strikingly larger than remainder. Inner row with,two scales and in addition two short stout spines which appear to take the positi,ons normally occupi,ed by scales. If we consider the,two pairs,of spines as equivalent to scales, the group ends up as part of Delamare s "multidentati group" to which it shows a number of similarities. Taken rom,one locality; Jisr esh Chouer, Syria, September 21, 1953. In swamp, Berlese funnel sample, wet soil mixed with dead reeds and roots. Mucronal formula (after Delamare) aa1a5. VARIATION Only four specimens were seen and in this series little variation was noted. In one specimen the unpaired teeth were reduced to,one extremely minute tooth on the first two pairs,of legs. The external row,of fringed scales were five in number on two specimens. DISCUSSION The presence of spines along with the fringed setae on the dens serves to separate this speci,es readily fr,om all previously described species of the genus. The presence of only two fringed scales,on the internal row in this species would appear to place the species in Delamar,e s Serroderus. As is pointed out elsewhere the limits of this taxon need re-definiti,on. EXPLANATION OF PLATE Fig. 1. Cyphoderus assimilis BSrner. Claw, specimen from Syria, approx. 500 x. Fig. 2. C. genneserae Carpenter. Clasp organ at base of dens, specimen from Lebanon, approx. 1200 x. Fig. 3. C. (Cyphoda) grassei Cassagnau and Delamare. Mucro and tip of dens typical Lebanese specimen, approx. 600 x. Fig. 4. C. (Cyphoda) grassei Cassagnau and Delamare. Basal dental organ, approx. 1200 x. Fig. 5. C. assimilis BSrner. Basal dental organ, specimen from Syria, approx. 1200 x. Fig. 6. C. (Cyphoda) grassei Cassagnau and Delamare. Semidiagrammatic representation of dorsal chaetotaxy. Fig. 7. C. genneserae Carpenter. As above, specimen from Lebanon. Fig. 8. C. spinatus n. sp. As above, paratype.

PsYcH, 1957 VoL. 64, PLATE 7 CttRISTIANSEN CYPI-IODERUS

84 Psyche [8eptembel" Cyphoderus assimilis BSrner Plate 6, figure 12, Plate. 7, figures 1, 5 Cyphoderus assimilis BSrner, 1906, Mitt. Naturhist. Mus. Hamburg 23:181. The true identity of this species is a difficult problem to solve. The species C. assimlilis, C. similis, C. subsimilis, and C. simulans all have in common" bidenticulate mucr.o, unguis without tunica and with two unpaired teeth,.only a single well developed basal ungual tooth and a clavate tenent hair. Although the absence.o the tunica, the unpaired median tooth, and clavate tenent appear to show some variation and much difference of interpretation, we have little choice other than to accept these as. real characteristics. It would appear from BSrner.s description that his original diagnosis is a.composite of two species. One of these is the species later described by Folsom as C. similis. The evidence for this is as 2.o.llows" First BSrner, 1906, described the species from two.collections; one from Egypt, and one from.orchids from the West Indies. Since the 2.orm Folsom described was from the West Indies, it. seems quite probable that it is equivalent to the latter form seen by BSrner. If this is accepted then a number of facts indicate that the.description is composite, and the Egyptian form is another species. BSrner says that the inner terminal scale is shorter to somewhat longer than the mucro. In the West Indian and Central American specimens it is never as long as the mucro. He says there is usually a small lamella upon the mucro; the West Indian and Central American material always have some lamella. This would mean that to account for his description the species seen by BSrner were two" one an Egyptian form sharing the.common characteristics of the group with a dental scale longer than mucro and without lamella on the mucro, and a second form later described by Folsom as C. similis. C. subsimilis o Delamare satisfies both requirements for the Egyptian species, but the figure of the unguis differs strikingly from that given by BSrner in 1913. I have collections from Syria which satisfy all of the characteristics listed above. The one major.disagreement between the species is the relative size of the

1957] Christiansen Collembola of Lebanon 85 mucrones. Since this, as most ratio distinctions, is very unstable it is not a serious objection. If this is accepted as the true assimilis then the following table can be used to separate the 2our species" assimilis? similis subsimilis simulans dental 2ormula 5/6 5/6 4/6 6/6 internal setae of 1 row tibiotarsus "smooth" "smooth"?? muero no lamella with lamella no lamella no lamella distal scale / mucro longer shorter longer shorter Distribution" Bahr Atibe, east of Damascus, Syria, July 10, 1953. Cyphodems albinus (Nicolet) Cyphoderus albinos (Nicolet), 1842, Nouv. Mere. Soc. Itelv. Sci. Nat. 6"1-88. Cassagnau and Delamare reported this from Dahr el Ain in Lebanon. I have two specimens taken from Latakia Syria, which appear to be this species. Unfortunately both were damaged in capture so that a certain identification is impossible. Cyphoderus bidenticulatus (Parona) Cyphoderus bidenticulatus (Parona), 1888, Ann. Mus. Civico Genova Ser. 6:83. Reported from two. caves in Lebano.n by Cassagnau and Delamare. Cyphoderus agnotus BSrner Cyphoderus agnotus BSrner, 1906, op. cir." 180-181. Although this. form has not been recorded from Lebanon.or Syria its widespread.occurrence throughout Palestine makes it quite probable that it does occur in the southern part of this area.

86 Psyche [september Genus Troglopedetes Cassagnau and Delamare recorded two new species of this cavernicolous genus from Lebanon (op. cir.). This known species of the genus, is largely circum-mediterranean in distribution; but, the existence of Paleotropical an,d Neotropical members indicates the possibility of a tr.opicopolitan but still undiscovered distribution. One new species was discovered in this work bringing the known Syrian-Lebanese forms to three. Troglopedetes orientalis Cassagnau & Delamare Troglopedetes orientalis, Cssagnau & Delmare, 1951, op. cit.: 385-387. These authors recorded this species from three localities in Lebanon. One record was non-cavernicolous. Troglopedetes vandeli Cassagnau & Delamare Troglopetes vandeli, Cassagnau & Delmre, 1951, op. cir.: 387-388. Taken from one cave in Lebanon. Trogopedetes canis n. sp. Plate 6, figur.es 6-8 Facies characteristic for the genus. Antennae with basal three segments subcylindrical and fourth segment divi.ded in middle and elongate fusiform in shape. Antennae covered with a variety,of setae of differ,ent types (see Plate 6, figure 6). Setae of Type A decreasing in size apically and with the ciliations becoming mor.e prominent. Setae of types F and B are limited to the fourth antennal segment while those of type C are most numerous here and upon a limited region on the inner margin of the apex of the second segment. Fourth segment with two scale-like knobs having a lateral axis, and a pair of small and.oval blades projecting out from this. In addition nine smooth setae are present. Scales.oval, with.extremely short fine striations. Dorsum of first thoracic segment with an anterior lateral fringe of large acuminate finely ciliate setae. Because of the dense covering of large scales, the remainder of the body setae are difficult to perceive, but the situation appears to be quite similar to that found in Cyphoderus. Dense with a single row of twenty-one spines. teeth. Mucro with four very small and two large dorsal

Christiansen-- Collembola of Lebanon 87 Tibiotarsi with all setae uniformly ciliate. Tenent hair prominent and clavate. Unguis with basal teeth w.ell developed, one definitely larger, two unpaired median teeth, the api.cal one being very minute. Median teeth more pr.ominent upon the hind unguis. Unguiculus acuminate and sparsely serrate along its posterior margin. Variation in dental spines: 22/19, 21/23, 22/21, 19/20, 19/23. Known only from the type locality: Dog River Cave, Lebanon, July, 1951 on Debris 1 km. inside cave. DISCUSSION This species is similar in many respects to T. orientalis but differ.s from this species in the shape.of the mucro, the number and disposition of the dental spines, possession of a clavate tenent hair, the type of setae upon the antennae (see below), and the structures of the ungues. In this last characteristic the present species ressembles T. machadoi Delamare, but it differs from this form in most respects, most strikingly in the relative length of the mucro. The setae upon the antennae of these animals furnish an excellent series of characteristics. Unfortunately only three species were availabl.e for comparison but as the following table will illustrate, these, indicated the value of further investigations.of this characteristic. In a.ddition to this the so-called sense organ.of the third antennal segment differs among the three forms (axis central in ruffoi and lateral in the other two). seta type orientalis ruffoi A * longer, ciliation more prominent B stalk longer than apical longer only slightly expansion expanded apically C unilaterally finely ciliate dense mass upon apex of third segment D * straight E * * F longer and definitely longer curved signifies similar to that illustrated for canis

88 Psyche [September Oncopodura ambigua n. sp. Plate 6, figures 1-5 Facies typical for genus. White, without trace of pigment or eyes. All antennal segments subovoid. Antennal segment four equipped with an apical conical projection and four blunt oval "sensory" setae. Remainder of setae of fourth segment slender smooth and acuminate. Third antennal segment with apical organ of two elliptical rods, with marked ladder-like striations, without clearly visible central axis. Remainder of antennae covered with a mixture of smooth setae (as in fourth antennal segment) and heavy conical finely striate setae. Postantennal organ of six lobes, the anterior pair lying at right angles to the long axis of the head, and the remainder radiating backward from this. Tibiotarsus without any clearly spatulate setae. Unguis untoothed, with a definite "triangular lamella" and an,o.pposing smaller but similar seta. Both structures are attached near the. base of the unguis. Empodial appendage simple, external edge straight and equipped with a short apical filament. Mucro slightly shorter than dens, equipped with an apical and three, dorsal teeth. A single large scale is attached to the base of the mucro. Dens clearly subdivided into two portions. Inner dorsal margin of dens equipped with four apically curved heavy ciliate setae. A sixth seta is on the,outer margin near the base of the dens. Remainder of dorsal surface with one large scale and five acuminate smooth setae. DISCUSSION In the series examined there was little variation outside of relative size. The large ciliate setae upon the inner margin may be five in number. Occasionally the basalmost of these setae is not curved at its apex. The P.A.O. is often sunken into the head and it is very difficult to puzzle out the exact arrangement o.f the lobes. The presence of a definite triangular ungual lamella combined with the presence of a six-lobed P.h.O. immediately serves to distinguish this species fr.om all those given in

t957] Christiansen- Collembola of Lebanon 89 Bonet (1943). In most characteristics this species would appear to be most closely allied to O. crassicornis Shoebotham and O. cruciata Boner, but it may readily be distinguished from both of these by the presence of the ungual lamella. This animal was extracted from a soil sample with a Berlese Funnel. Since only one locality is known, the range of the animal cannot be determined. Known.only rom the type locality" Campus American University, Beirut, Lebanon, various dates, 1953. LITERATURE CITED (See also Bibliography to Part I) BONET, M. F. 1943. Sobre La Classification de los Oncopoduridae Anales de la Escuela Nac. Cien. Biol. 3 (1-2):127-153. CHRISTIANSEN, K. 1956. The Collembola of Lebanon and Western Syria. Part I. General Considerations and the Family Onychiuridae. Psyche 63 (4): 119-133. DELAMARE-DEBOUTTEVILLE CL. 1948. Collemboles Termitophiles et Myrmecophiles. Arch. Zool. Experimentale Gen. 85:261-425.

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